Of Peru: a Survey of the Families

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Faculty Publications: Department of Entomology Entomology, Department of

2015 Beetles (Coleoptera) of Peru: A Survey of the Families. Scarabaeoidea Brett .C Ratcliffe University of Nebraska-Lincoln, [email protected]

M. L. Jameson Wichita State University, [email protected]

L. Figueroa Museo de Historia Natural de la UNMSM, [email protected]

R. D. Cave University of Florida, [email protected]

M. J. Paulsen University of Nebraska State Museum, [email protected]

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Ratcliffe, Brett .;C Jameson, M. L.; Figueroa, L.; Cave, R. D.; Paulsen, M. J.; Cano, Enio B.; Beza-Beza, C.; Jimenez-Ferbans, L.; and Reyes-Castillo, P., "Beetles (Coleoptera) of Peru: A Survey of the Families. Scarabaeoidea" (2015). Faculty Publications: Department of Entomology. 483. http://digitalcommons.unl.edu/entomologyfacpub/483

This Article is brought to you for free and open access by the Entomology, Department of at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Faculty Publications: Department of Entomology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Authors Brett .C Ratcliffe, M. L. Jameson, L. Figueroa, R. D. Cave, M. J. Paulsen, Enio B. Cano, C. Beza-Beza, L. Jimenez-Ferbans, and P. Reyes-Castillo

This article is available at DigitalCommons@University of Nebraska - Lincoln: http://digitalcommons.unl.edu/entomologyfacpub/ 483 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY 88(2), 2015, pp. 186–207 Beetles (Coleoptera) of Peru: A Survey of the Families. Scarabaeoidea

1,10 2 3 4 5 B. C. RATCLIFFE, M. L. JAMESON, L. FIGUEROA, R. D. CAVE, M. J. PAULSEN, 6 7 8 9 ENIO B. CANO, C. BEZA-BEZA, L. JIMENEZ-FERBANS, AND P. REYES-CASTILLO

ABSTRACT: The 1042 species of Scarabaeoidea known to occur in Peru are listed with their taxonomic placement in families, subfamilies, and tribes.

Peru is known for high species richness, endemism, and habitat complexity, but despite this interest in Peruvian biodiversity, little is known about the beetle fauna of the country (Larsen et al., 2011). The following checklist reflects our most current knowledge about the composition of the superfamily Scarabaeoidea in Peru. Compared with the neighboring country of Chile in which 247 species and 66 genera of scarabaeoid beetles are recorded (Elgueta, 2000), we record 1045 species of Peruvian scarabaeoid beetles, but we know the number will increase with additional exploration and discovery. This checklist was assembled by compiling data from print and online catalogs, the primary literature that describes new species or reviews genera, some research collections, and personal communications from some entomologists who have personally conducted research in Peru. Clearly, there are other species of scarabaeoids that are not yet recorded from Peru because their data have never been included in the literature. Conversely, there may be a few species recorded from Peru in the older catalogs that do not actually occur in Peru because they were incorrectly identified. Moreover, disparate levels of knowledge exist between scarab groups because some have undergone considerable study while others have not. In the lists below, an asterisk (*) indicates a species that is known only from Peru. Based upon our data, there are large numbers of scarabs endemic to Peru. This suggests that the scarab beetle fauna of Peru is amazingly rich, and that further exploration and discovery will reveal many more species. Scarab beetles are an important component of Peruvian biodiversity because they are important pollinators of plants, decomposers of waste, economically important pests, and some are valuable culturally for adornment or as a food source. Knowledge about

1 Systematics Research Collections, University of Nebraska, Lincoln, Nebraska 68588-0514, USA; e-mail: [email protected]. 2 Department of Biological Sciences, Wichita State University, Wichita, Kansas 67260-0026, USA; e-mail: [email protected]. 3 Departmento de Entomologı´a, Museo de Historia Natural de la UNMSM, Lima, PERU; e-mail: [email protected]. 4 Indian River Research and Education Center, University of Florida, Fort Pierce, Florida 34945-3138, USA; e-mail: [email protected]. 5 Systematics Research Collections, University of Nebraska, Lincoln, Nebraska 68588-0514, USA; e-mail: [email protected]. 6 Systematic Entomology Laboratory, Universidad del Valle de Guatemala, GUATEMALA; e-mail: [email protected]. 7 Department of Biological Sciences, University of Memphis, Memphis, Tennessee 38152, USA; e-mail: [email protected]. 8 Universidad de La Guajira, Riohacha, COLOMBIA; e-mail: [email protected]. 9 Instituto de Ecologı´a, A. C., Xalapa 91070, Veracruz, MEXICO; e-mail: [email protected]. 10 Corresponding Author E-mail: [email protected]. Accepted 21 April 2014; Revised 1 December 2014 E 2015 Kansas Entomological Society VOLUME 88, ISSUE 2 187

Peruvian biodiversity is pivotal for conserving and managing natural resources, food security, poverty reduction, health, biosecurity, new industrial product development, and ecotourism (Smith et al., 2011).

Geotrupidae Diversity in Peru: 1 subfamily, 4 genera, and 11 species. Recognition: The body shape is oval or round, and the head is not deflexed. The antennae are 11-segmented with a 3-segmented, opposable club with all antennomeres tomentose. The eyes are completely or partially divided by a canthus. The clypeus is often with a tubercle or horn. The labrum is truncate, prominent, and produced beyond the apex of the clypeus. The mandibles are prominent and produced beyond the apex of the labrum. The pronotum is convex with a base wider than or subequal to the elytral base and with or without tubercles, ridges, horns, or sulci. The elytra are convex, with or without striae. The pygidium is concealed by the elytra (Jameson, 2002a). Habitat: Life histories of the geotrupids are diverse, and food habits vary from saprophagous to coprophagous and mycetophagous. Adults of most species are secretive, living most of their life in burrows. Although adults do not tend larvae, adults provision food for larvae in brood burrows. Adults dig vertical burrows (15– 200 cm in depth) and provision larval cells with dead leaves, cow dung, horse dung, or humus. Burrows of some species extend to a depth of 3.0 m (Jameson, 2002a). Notes: The family Geotrupidae includes 68 genera and about 620 species (Scholtz and Browne, 1996). The subfamily Geotrupinae does not occur in South America. The following checklist of Peruvian Scarabaeidae is from Howden (1985, 2002) and Howden and Martı´nez (1978).

BOLBOCERATINAE Athyreini Athyreus bicornus Howden, 2002* Neoathyreus fallolobus Howden, 2006 Athyreus larseni Howden, 2002* Neoathyreus ornatus Howden, 1985* Athyreus martinezi Howden, 1995 Neoathyreus rufobrunneus Howden, 1985 Athyreus pyriformis Howden and Martı´nez, 1978* Neoathyreus rufoventris Howden, 1985* Athyreus tribuliformis Felsche, 1909*

Bolboceratini Bolboceras baeri Boucomont, 1902 Zefevazia peruana (Boucomont, 1902)* Lucanidae Diversity in Peru: 1 subfamily, 14 genera, 40 species. Recognition: The head is prognathous and not deflexed. The antennae are geniculate or straight, 10-segmented, and with a 3–7 segmented club (all antennomeres unopposable and tomentose). The first antennomere is often subequal in length to the remaining antennomeres. Habitat: Lucanids are usually associated with decaying wood and logs in coniferous and deciduous forests. Adults of some species are attracted to lights at night and some feed at sap flows from fluxing trees. The eggs are usually laid in crevices in bark or logs, and the larvae feed on decaying wood. Notes: The stag beetle family Lucanidae includes four subfamilies, 108 genera, and around 1,500 species worldwide. In the New World, there are 41 genera and 223 188 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY species with representatives of each of the four subfamilies (Paulsen and Ratcliffe, 2005). Most New World taxa are Neotropical. The following checklist of Peruvian Lucanidae is from Paulsen (2013).

LUCANINAE Brasilucanini Brasilucanus acomus Ratcliffe, 1984

Chiasognathini Sphaenognathus alticollis Mo¨llenkamp, 1912 Sphaenognathus peruvianus (Waterhouse, 1869) Sphaenognathus gaujoni (Oberthu¨r, 1885) Sphaenognathus prionoides Buquet, 1838 Sphaenognathus giganteus Boileau, 1911 Sphaenognathus xerophilus Bartolozzi and Onore, Sphaenognathus monguilloni Lacroix, 1972* 2006*

Sclerostomini Aegognathus aguirei Arnaud and Bomans, 2007* Incadorcus damasoi Arnaud and Bomans, 2006* Aegognathus confusus Arnaud and Bomans, 2006* Incadorcus michelleae Arnaud and Bomans, 2006* Aegognathus leuthneri leuthneri Van der Poll, 1886 Incadorcus shaunai Arnaud et al., 2007* Aegognathus leuthneri damasoi Arnaud and Incadorcus zugeri Arnaud and Bomans, 2006* Bomans, 2006* Metadorcinus beneshi (Martı´nez, 1953) Aegognathus similis Arnaud and Bomans, 2006* Metadorcinus lineatus (Deyrolle, 1864)* Aegognathus soulai Arnaud and Bomans, 2004* Metadorcinus yamauchii Arnaud et al., 2008* Aegognathus waterhousei Leuthner, 1883* Metadorcus ebeninus Deyrolle, 1864 Andinolucanus inesae Arnaud and Bomans, 2006* Onorelucanus boileaui (Weinreich, 1960)* Arnaudius bomansi Grossi and Bartolozzi, 2011* Onorelucanus marujae (Arnaud et al., 2008)* Arnaudius digennaroi (Arnaud et al., 2008)* Onorelucanus noguchii (Arnaud and Bomans, Arnaudius koikei (Arnaud et al., 2007)* 2007)* Auxicerus platyceps Waterhouse, 1883* Pseudoscortizus incredibilis Arnaud et al., 2008 Cantharolethrus azambrei Boileau, 1897 Sclerostomus bartolozzii Arnaud and Bomans, Cantharolethrus elongatus Lacroix, 1982* 2007* Cantharolethrus steinheili Parry, 1875* Sclerostomus damasoi Arnaud and Bomans, 2006* Incadorcus ashaninka Grossi, 2011* Sclerostomus wendyae Arnaud and Bomans, 2006* Incadorcus cuzcoensis Arnaud et al., 2007* Scortizus prodigiosus Arnaud et al., 2007*

Passalidae Diversity in Peru: 1 subfamily, 2 tribes, 6 genera, 50 species. Recognition: The body shape is elongate-cylindrical and depressed, and the length is 13–80 mm. Their color is black and shiny. The head is prognathous, narrower than the thorax, and often with a dorsomedian horn. The antennae have ten antennomeres including a 3 to 5-segmented club that is not opposable and not geniculate but is capable of being rolled together. The mandibles are large and project beyond the apex of the labrum, and the apex of the labrum is large, rounded, and toothed. The pronotum is broader than the head, quadrate, and the surface is smooth with a median, longitudinal groove. The elytra are elongate, parallel sided, with rounded apices and well-developed striae. The pygidium is concealed by the elytra (Schuster, 2002). Habitat: Passalid adults live in well-decayed logs and stumps with their larvae in subsocial family groups. All stages are found in galleries of wood that are excavated by the adults. Eggs are usually placed together in a nest of frass. Adults and larvae communicate by stridulating and can produce different calls. Adults care for larvae and prepare food by chewing it and presumably mixing it with saliva (Schuster, 2002). One species (Ptichopus angulatus [L.]) inhabits the waste chamber in nests of leaf cutter ants, Atta species. VOLUME 88, ISSUE 2 189

Notes: Most New World taxa occur in the Neotropics. Hincks and Dibb (1935) cited passalid species from Peru, and Van Doesburg (1942) provided a checklist of Peruvian Passalidae.

PASSALINAE

Passalini Passalus abortivus Percheron, 1835 Passalus plicatus Percheron, 1835 Passalus aduncus Erichson, 1847 Passalus prominens Gravely, 1918* Passalus anguliferus Percheron, 1835 Passalus pubicostatus (Kuwert, 1898) Passalus arrowi Hincks, 1934 Passalus pugionatus Burmeister, 1847 Passalus barrus Boucher and Reyes-Castillo, 1991 Passalus pugionifer (Kuwert, 1891)* Passalus caelatus Erichson, 1847 Passalus punctiger Le Peletier and Servillle, 1825 Passalus coniferus Eschscholtz, 1829 Passalus rhodocanthopoides (Kuwert, 1891) Passalus convexus Dalman, 1817 Passalus rotundatus Hincks, 1940* Passalus elfriedae Lu¨derwaldt, 1931 Passalus rusticus Percheron, 1835 Passalus episcopus (Kuwert, 1898)* Passalus schneideri (Kuwert, 1898)* Passalus huebneri (Kuwert, 1898)* Passalus spinifer Percheron, 1835 Passalus inca (Zang, 1905)* Passalus zangi Hincks, 1934 Passalus interruptus (L., 1758) Paxillus camerani (Rosmini, 1902) Passalus interstitialis Eschscholtz, 1829 Paxillus.forsteri Lu¨derwaldt, 1927 Passalus latifrons Percheron, 1841 Paxillus leachi MacLeay, 1819 Passalus occipitalis Eschscholtz, 1829 Spasalus crenatus (MacLeay, 1819) Passalus peruvianus (Kuwert, 1898) Spasalus kaupi Boucher, 2004*

Proculini Popilius amazonicus Gravely, 1918* Veturius inca Boucher, 2006 Popilius marginatus (Percheron, 1835) Veturius libericornis Kuwert 1891 Verres furcilabris (Eschscholtz, 1829) Veturius sinuosus (Drapiez, 1820) Veturius amazonicus Boucher, 2006 Veturius spinipes (Zang, 1905) Veturius arawak Boucher, 2006 Veturius standfussi Kuwert, 1891 Veturius cephalotes (Le Peletier and Serville, 1825) Veturius tarsipes Boucher, 2006* Veturius ecuadoris Kuwert, 1898 Veturius unicornis Gravely, 1918 Veturius guntheri Kuwert, 1898 [5 peruvianus of Veturius yahua Boucher, 2006 Arrow]

Trogidae Diversity in Peru: 1 subfamily, 2 genera, 10 species. Recognition: Adult trogids are recognized by their warty, brown to gray to black, dirt-encrusted appearance and flat abdominal sternites. Habitat: Adults and larvae are among the last scavengers that visit the dry remains of dead animals, where they feed on feathers, fur, and skin. They also feed on organic matter found in the nests of mammals and birds. Notes: The world fauna consists of 4 genera and 300 species (Scholtz, 1982). The following checklist of Peruvian trogids is from Scholtz (1982).

TROGINAE Omorgus howelli (Howden and Vaurie, 1957) Polynoncus bullatus (Curtis, 1845) Omorgus suberosus (Fabricius, 1775) Polynoncus ecuadorensis (Vaurie, 1962) Omorgus persuberosus (Vaurie, 1962) Polynoncus gordoni (Steiner, 1981)* Polynoncus aeger (Gue´rin-Me´neville, 1844)* Polynoncus gemmingeri (Harold, 1872) Polynoncus aricensis (Gutie´rrez, 1950) Polynoncus peruanus (Erichson, 1847) Polynoncus brevicollis (Eschscholtz, 1822) Polynoncus sallei (Harold, 1872) 190 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Ochodaeidae Diversity in Peru: 1 subfamily, 1 genus, 1 species. Recognition: Ochodaeids are elongate and convex. Their color is testaceous, brown, reddish brown, brown, black, or occasionally bicolorous. The head is not deflexed. The antennae have 9 or 10 antennomeres, with a 3-segmented, opposable club, and all antennomeres are tomentose. The clypeus is simple or tuberculate on the anterior margin. The labrum is produced beyond the apex of the clypeus, often bilobed and emarginate, and prominent. The prominent mandibles are produced beyond the apex of the labrum. The pronotum is convex, subquadrate, mostly punctate and setose and without tubercles, ridges, horns, or sulci. The mesotibia at the apex has at least one apical spur pectinate. The elytra are convex, with or without striae, often punctate, or granulate and setose, but some are smooth. The pygidium is exposed or concealed by the elytra (Carlson, 2002). Habitat: Ochodaeidae are most often collected at lights, sometimes in large numbers. Adults of a few species are active during the day. Many species prefer sandy areas. Adults may spend the day in subterranean burrows, where they may feed on fungi. Little is known about the biology of Ochodaeidae. There are few observations of adult or larval habits except that adults of most species are nocturnal and are attracted to lights (Carlson, 2002). Notes: The family Ochodaeidae includes 10 genera and about 80 species worldwide (Carlson and Paulsen, 2012).

OCHODAEINAE Parochodaeus bituberculatus (Erichson, 1847)*

Hybosoridae Diversity in Peru: 3 subfamilies, 11 genera, 33 species. Recognition: Hybosorids are light brown to black and glossy. The head is not deflexed. The antennae are 10-segmented with a 3-segmented, opposable club with the last 2 antennomeres tomentose. The first antennomere of the club is hollowed to receive antennomeres 2 and 3. The clypeus generally lacks a tubercle or horn. The prominent labrum is truncate and produced beyond the apex of the clypeus. The large mandibles project beyond the apex of the labrum and have the external edge rounded with the apices pointed. Habitat: Little life history information is known for hybosorids. Adults feed on both invertebrate and vertebrate carrion in the early stages of decomposition. Some species are found in dung, and others are attracted to lights at night (Jameson, 2002b). Ceratocanthines are commonly found in clusters of dead leaves and can be collected using a beating sheet. Notes: The family Hybosoridae worldwide contains 4 subfamilies, 35 genera, and about 220 species (Ocampo, 2006; Ocampo and Ballerio, 2006). Species are widely distributed in the tropics. The classification below is from Ocampo and Ballerio (2006).

ANAIDINAE Anaides onofrii Ocampo, 2006 Chaetodus allsoppi Martı´nez, 1988* Anaides rugosus Robinson, 1948* Chaetodus asuai Martı´nez, 1956 VOLUME 88, ISSUE 2 191

Chaetodus mimi Ocampo, 2006* Hybochaetodus erugocarinatus Ocampo, 2006* Chaetodus smithi Ocampo, 2006* Hybochaetodus flaco Ocampo, 2002* Chaetodus tricarinatus Ocampo, 2006 Hybochaetodus obscurus Arrow, 1909* Hybochaetodus disruptus Ocampo, 2006*

CERATOCANTHINAE Anopsiostes punctatus Paulian, 1982 Ceratocanthus suturalis (Lansberge, 1887) Astaenomoechus americanus (Boucomont, 1936) Germarostes antiquus (Erichson, 1843) Astaenomoechus criberrimus Paulian, 1982* Germarostes aphodioides (Illiger, 1800) Ceratocanthoides undatus (Petrovitz, 1973) Germarostes carltoni Howden and Gill, 2005* Ceratocanthus clypealis (Lansberge, 1887)* Germarostes geayi Paulian, 1982* Ceratocanthus inca Paulian, 1982* Germarostes macleayi (Perty, 1830) Ceratocanthus mathani Paulian, 1982* Germarostes semituberculatus (Germar, 1843) Ceratocanthus perpunctatus Paulian, 1982* Germarostes sulcipennis Harold, 1875* Ceratocanthus politus (Erichson, 1843) Trachycrusus lescheni Howden and Gill, 1995* Ceratocanthus punctulatus Lansberge, 1887* Trachycrusus striatulus Howden and Gill, 1995* Ceratocanthus seriatus (Erichson, 1843)

HYBOSORINAE Coilodes punctipennis Arrow, 1909

Scarabaeidae Diversity in Peru: 10 subfamilies, 194 genera, and 898 species. Recognition: Scarabs are 2.0–160.0 mm in length with variable shape and color, with or without metallic reflections, and with or without vestiture. The antennae have 10 antennomeres (some 7–12) with a 3 to 5-segmented, opposable club; the club has the apical antennomeres nearly glabrous (Melolonthinae, Dynastinae, Rutelinae, Cetoniinae) or with all antennomeres tomentose (Aphodiinae, Scarabaeinae). The clypeus is with or without a tubercle or horn. The labrum is distinct in most taxa, and produced beyond the apex of the clypeus or not. The mandibles are variable, produced beyond the apex of the labrum or not. The pronotum is variable, with or without horns or tubercles. The elytra are convex or flattened and with or without striae. The pygidium is concealed by the elytra (Aphodiinae, Scarabaeinae) or exposed (Scarabaeinae, Melolonthinae, Dynasti- nae, Rutelinae, Cetoniinae). The scutellum is exposed or not, and its shape is triangular or parabolic. The legs have transverse or conical coxae; the protibiae are tridentate, bidentate, or serrate on the outer margin; the meso- and metatibia at the apex have 1 or 2 spurs; the tarsi are 5-5-5, and the anterior tarsi are absent in some Scarabaeinae; the tarsal claws are variable, equal in size or not, and simple or toothed. The abdomen has 6 free sternites; the 7 functional abdominal spiracles are situated in the pleural membrane (Aphodiinae, Scarabaeinae) or in the sternites and tergites (Melolonthinae, Dynastinae, Rutelinae, and Cetoniinae). Habitat: Scarab beetles occupy many habitats ranging from deserts to tropical rainforests and have diverse life histories. Depending on the group, adults feed on foliage, dung, flowers, or sap while the larvae are phytophagous, coprophagous, or detritivorous. Some are diurnal, while many are nocturnal in their activity patterns. Notes: Members of the subfamily Aphodiinae, known as the aphodiine dung beetles, are primarily dung and detritus feeders. All of the species are small, rarely exceeding 12 mm in length. Worldwide, the subfamily contains 12 tribes, 192 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY approximately 280 genera, and 3,200 species with 9 tribes, 128 genera and 816 species in the New World (Skelley, 2008). In Peru, there are 24 genera and 63 species (Skelley, 2008). The subfamily Scarabaeinae is commonly referred to as dung beetles. Most species feed on mammal dung, but others specialize upon the dung of other vertebrates and invertebrates as well as on carrion, mushrooms, rotting fruit, and other decomposing plant material. The world fauna includes over 5,000 described species in 234 genera, with about 1,800 species just in the genus Onthophagus (Gill, 2005). The tribal classification below follows that of Hanski and Cambefort (1991). Peru is incredibly rich in dung beetles, and over 150 species are recorded from a single lowland forest site in Madre de Dios (Los Amigos, Peru) (Trond Larsen, personal communication to BCR, February 2014). Larsen and Geńier (2008a, b) provided color guides for identifying dung beetles at two different biological stations in Peru. Peru has 40 genera and 278 species, but many more will be recognized in the future, especially in the genera Ateuchus, Canthidium, Dichotomius, and Uroxys. In comparison, neighboring Brazil includes 49 genera and 618 species (Vaz-de-Mello, 2000). The Melolonthinae, often called May beetles, June beetles, or leaf chafers, are cosmopolitan in distribution and one of the largest and most diverse subfamilies of Scarabaeidae. There are approximately 750 genera and over 11,000 species worldwide, with about 90 genera in the New World (Evans and Smith, 2005). Peru has 20 genera and 82 species, but this number will increase as more species are recognized. This is not now easily accomplished because identification of some genera and many species remains in a state of confusion. The tribes are still unresolved due to the lack of definition and inconsistent use of characters (Evans and Smith, 2005). Reliable identifications are difficult since the majority of descriptions prior to 1940 lacked illustrations or comparisons with other similar species. The subfamily Rutelinae (leaf chafers) includes 7 tribes, 2 of which do not occur in Peru (Adoretini from the Old World and Alvarengiini from southern Brazil), and approximately 4,300 species. Based on this checklist, the leaf chafers of Peru encompass a rich fauna with 52 genera, 270 species and subspecies, and 150 endemic species. The neighboring country of Ecuador includes 53 genera, 298 species, and 92 endemic species (Paucar-Cabrera, 2005). The last compilation of ruteline species of Peru was a product of the Hamburg Southern Peru Expedition in 1936, and this yielded 29 genera and 85 species (Ohaus, 1952). Because of the economic importance and interest in the Rutelinae, classification and nomenclature of the group is undergoing extensive revisions. This checklist reflects changes in the tribes Anomalini (Ramı´rez-Ponce and Moroń, 2009; Moroń and Ramı´rez-Ponce, 2012), Anoplognathini (Smith, 2003), Anatistini (previously Spodochlamyini; Jameson and Ratcliffe, 2011), Geniatini (Hawkins and Jameson, 2005; Soula, 2010), and Rutelini (Soula, 1998, 2002a, b, 2003, 2005, 2006a, b, 2008, 2009, 2010, 2011a, b; Jameson, 1998; Moore and Jameson, 2013). It should be noted that Soula’s species concept was typological as well as topological; specimens that displayed variation from the type specimen and/or specimens from unique localities were often described as new species, subspecies, or varieties, thus reflecting a “stamp collector” view of biodiversity. As a result, Soula greatly overestimated ruteline diversity by describing many new genera and species that we believe will become VOLUME 88, ISSUE 2 193 invalid. In this checklist, Soula’s taxa comprise nearly half of the ruteline diversity (119 species and 31 subspecies) in Peru. Because Soula’s names were established in accordance with the rules of zoological nomenclature, the names must be considered as available. We do not address validity of Soula’s species and genera here, but we caution the user in accepting Soula’s classification and species concepts. Because of Soula’s overestimation of ruteline diversity, lack of survey throughout South America, and poor taxonomic foundation, the Rutelinae are currently poor bioindicators for habitat conservation. Rutelines are important ecologically as pollinators and decomposers; agriculturally as consumers of crop leaves, roots, and fruits; and culturally as food and decorative adornment. Species in the subfamily are often highly metallic and large (up to 50 mm in length). Because of their beauty and abundance, the Shuar people (also Achuar, apach, inkis, jı´varos, or jı´baros) of the eastern Amazonian areas in Peru and Ecuador use the bodies of some rutelines (especially Chrysophora chrysochlora) to create headpieces and necklaces (Ratcliffe, 2006). These decorative pieces are now popular among tourists and local people alike. The subfamily Dynastinae is one of the most conspicuous subfamilies of the beetle family Scarabaeidae. Members of the subfamily occur in all the major biogeographic areas of the world (except the polar regions), although most species are found in the tropics, specifically the New World tropics. Dynastines comprise 196 genera and about 1,500 species grouped among 8 tribes. Six tribes, 87 genera, and at least 800 species occur in the New World, and most of these species occur in the Neotropics. Our checklist for Peru currently has 39 genera and 182 species. By comparison, Mexico, another large and megadiverse country, has 30 genera and 196 species (Ratcliffe et al., 2013), while Costa Rica and Panama together have 36 genera and 158 species (Ratcliffe, 2003). Adult dynastines are small (4 mm) to very large (160 mm) beetles. The males in some species (principally Dynastini, Agaocephalini, and Oryctini) possess prominent and often spectacular horns on the head and/or prothorax which, together with their large size, have given rise to such popular names for them as “rhinoceros,” “elephant”, “hercules”, and “unicorn” beetles. In fact, the entire subfamily is usually referred to as the rhinoceros beetles even though the majority of species do not possess horns. The adults of nearly all species are nocturnal or crepuscular, and many are readily attracted to lights at night. Adult dynastines are known to feed on ripe or rotting fruits, slime fluxes, and plant roots. The adults of some Cyclocephalini are important pollinators of palms and aroids when they feed on the floral parts of these plants. The subfamily Cetoniinae, or flower chafers, are most abundant in Africa and Asia. In the New World they comprise 6 tribes in 41 genera with approximately 275 species (Krajcik, 2012). In Peru there are 4 tribes containing 8 genera and 18 species. The genus Gymnetis is still in need of revision, and the species names below have presumed synonyms that may turn out to be valid species. The following checklist of Peruvian Scarabaeidae is extracted from many sources, principal among them are Ocampo and Colby (2009) (Allidiostomatinae); Skelley (2008), Smith and Skelley (2007), and Stebnicka (2005, 2007, 2009) (Aphodiinae); Edmonds (1994, 2000), Edmonds and Zidek (2010, 2012), and Figueroa et al. (2014) (Scarabaeinae in part); Evans and Smith (2009) (Melolonthinae); Machatschke (1972), Villatoro (2002), Jameson (1998), and Soula (1998–2011) (Rutelinae); Colby 194 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

(2009) (Orphninae), Endro¨di (1966–1985) (Dynastinae); Erichson (1847) (Peru); Blackwelder (1944) (Scarabaeoidea), and Krajcik (2012). The older checklists in particular should be used with caution because classification and some names may have changed.

APHODIINAE

Aphodiini Aidophus infuscatopennis (Schmidt, 1909) Neotrichaphodioides woytkowskii Dellacasa, Dellacase, Aphodius pseudolividus Balthasar, 1941 and Skelley, 2010* Blackburneus laxepunctatus (Schmidt, 1910) Orodaliscoides rugosiceps (Harold, 1859) Gonaphodiellus castanescens (Petrovitz, 1973) Paranimbus peruanus (Erichson, 1834) Gonaphodiellus chapini (Hinton, 1934) Trichaphodiellus brasiliensis (Laporte, 1840) Gonaphodiellus nigrinus (Schmidt, 1916)*

Eupariini Ataeniopsis regulis (Balthasar, 1947) Ataenius petrovitzi Balthasar, 1960 Ataenius abancay Stebnicka, 2005* Ataenius picinus Harold, 1868 Ataenius aequatorialis Petrovitz, 1961 Ataenius platensis (Blanchard, 1846) Ataenius atramentarius (Erichson, 1847) Ataenius rubrotessellatus (Blanchard, 1843) Ataenius attenuator Harold, 1874 Ataenius santarosae Stebnicka, 2007 Ataenius buenavistae Stebnicka, 2001 Ataenius sculptilis Harold, 1868 Ataenius carinator Harold, 1874 Ataenius siminasus Petrovitz, 1973 Ataenius catenulatus (Erichson, 1847)* Ataenius strigicaudus Bates, 1887 Ataenius columbicus Harold, 1889 Ataenius tambopatae Stebnicka, 2001* Auperia andamanensis (Koshantschikov, 1916) Ataenius complicatus Harold, 1869 Auperia capitosus (Harold, 1867) Ataenius costulifer Balthasar, 1941* Auperia huebneri (Petrovitz, 1970) Ataenius crenaticollis Petrovitz, 1973 Auperia iquitosae (Stebnicka, 2002) Ataenius gracilis (Melsheimer, 1844) Auperia loretoensis (Stebnicka, 2002)* Ataenius huanus Stebnicka, 2007 Euparixia boliviana Gordon and McCleve, 2003 Ataenius icanus Balthasar, 1951* Euparixiodes johnsoni Stebnicka, 1998 Ataenius impiger Schmdt, 1916 Lomanoxoides mapitunari Stebnicka and Skelley, Ataenius lamarensis Stebnicka, 2007 2005* Ataenius montanus Schmidt, 1911 Odontolytes iquitosae (Stebnicka, 2007) Ataenius morator Harold, 1869 Passaliolla corticalis (Bates, 1887) Ataenius napoensis Stebnicka, 2007 Saprosites dentipes Harold, 1867 Ataenius nugator Harold, 1880 Saprosites parallelus Harold, 1867 Ataenius palmaritoensis Stebnicka, 2007 Saprosites sulcatus Harold, 1869

Odontolochini Amerilochus cinereus Skelley, 2007* Saprositellus denticulatus Balthasar, 1967 Saprolochus tambopatae Stebnicka and Galante, Saprositellus peruanus Stebnicka, 2003* 2007 Stebnickiella zosterixys Skelley, 2007* Saprolochus tridentatus Skelley, 2007

Psammodiini Mysarus peruanus Petrovitz, 1962*

Rhyparini Aschnarhyparus peregrinus (Hinton, 1934)

Stereomerini Termitaxis holmgreni Krikken, 1970* VOLUME 88, ISSUE 2 195

SCARABAEINAE

Onthophagini Digitonthophagus gazella (Fabricius, 1787) Onthophagus osculatii Gue´rin-Me´neville, 1855 Onthophagus bidentatus Drapiez, 1819* Onthophagus ptox Erichson, 1847* Onthophagus clypeatus Blanchard, 1846 Onthophagus ranunculus Arrow, 1913 Onthophagus coscineus Bates, 1887 Onthophagus rhinophyllus Harold, 1868 Onthophagus haematopus Harold, 1875 Onthophagus rubrescens Blanchard, 1846 Onthophagus marginicollis Harold, 1880 Onthophagus schunckei Paulian, 1936* Onthophagus onorei Zunino and Halffter, 1998 Onthophagus sp. aff. tristis Harold, 1873 Onthophagus onthochromus Arrow, 1913 Onthophagus xanthomerus Bates, 1887 Onthophagus ophion Erichson, 1847 Onthophagus sp. aff. xanthomerus Bates, 1887

Oniticellini Eurysternus caribaeus (Herbst, 1789) Eurysternus lanuginosus Geńier, 2009 Eurysternus cayennensis Laporte, 1840 Eurysternus marmoreus Laportee, 1840 Eurysternus contractus Geńier, 2009 Eurysternus nigrovirens Geńier, 2009 Eurysternus foedus Gue´rin-Meńeville, 1830 Eurysternus plebejus Harold, 1880 Eurysternus gracilis Geńier, 2009 Eurysternus squamosus Geńier, 2009 Eurysternus hamaticollis Balthasar, 1939 Eurysternus streblus Genier, 2009 Eurysternus howdeni Geńier, 2009 Eurysternus strigilatus Geńier, 2009 Eurysternus hypocrita Balthasar, 1939 Eurysternus vastiorum Martıńez, 1988 Eurysternus inca Geńier, 2009 Eurysternus wittmerorum Martıńez, 1988 Eurysternus inflexus Germar, 1824

Canthonini Anisocanthon villosus (Harold, 1868) Canthon septemmaculatus septemmaculatus Canthon aberrans (Harold, 1868) (Latreille, 1807) Canthon aequinoctialis Harold, 1868 Canthon septemmaculatus histrio LePeletier and Canthon angustatus Harold, 1867 Serville, 1828 Canthon balteatus Boheman, 1858 Canthon sericatus Schmidt, 1922 Canthon bimaculatus Schmidt, 1922 Canthon simulans Martıńez, 1950 Canthon brunneus Schmidt, 1922 Canthon smaragdulus (Fabricius, 1781) Canthon chiriguano Martinez and Halffter, 1972 Canthon subhyalinus Harold, 1867 Canthon coloratus Schmidt, 1922 Canthon triangulare Drury, 1870 Canthon conformis Harold, 1868 Canthon unicolor Blanchard, 1843 Canthon fulgidus Redtenbacher, 1867 Canthon velutinus Harold, 1868 Canthon fuscipes Erichson, 1847 Canthon virens chalybaeus Blanchard, 1843 Canthon gemellatus Erichson, 1847 Canthonella sp. aff. amazonica Ratcliffe and Smith, Canthon helleri Schmidt, 1922 1999 Canthon janthinus Blanchard, 1843 Canthonella barrerai Halffter and Martıńez, 1968 Canthon juvencus Harold, 1868 Canthonella sp. aff. catharinensis Pereira and Canthon laesum Erichson, 1847 Martıńez, 1956 Canthon lituratus Germar, (1813) Canthonella cf. gomezi Halffter and Martıńez, 1968 Canthon luteicollis Erichson, 1847 Canthonidia rubromaculata (Blanchard, 1846) Canthon sp. aff. matthewsi Martıńez and Halffter, Cryptocanthon campbellorum Howden, 1973 1972 Deltochilum aequinoctiale Buquet, 1844 Canthon monilifer Blanchard, 1843 Deltochilum amazonicum Bates, 1887 Canthon mutabilis Lucas, 1857 Deltochilum cf. aureopilosum Paulian, 1939 Canthon muticus Harold, 1867 Deltochilum burmeisteri Harold, 1867 Canthon pallidus Schmidt, 1922 Deltochilum carinatum Westwood, 1837 Canthon sp. aff. pallidus Schmidt, 1922 Deltochilum crenulipes Paulian, 1938 Canthon paraguayanum Balthasar, 1939 Deltochilum erodioides Harold, 1867 Canthon quadriguttatus (Olivier, 1789) Deltochilum fuscocupreum Bates, 1870 Canthon quinquemaculatus Laporte, 1840 Deltochilum granulatum Bates, 1870 Canthon rubrescens Blanchard, 1843 Deltochilum howdeni Martıńez, 1955 Canthon semiopacus Harold, 1868 Deltochilum hypponum Buquet, 1844 196 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Deltochilum sp. aff. komareki Balthasar, 1939 Scybalocanthon aereus (Schmidt, 1922) Deltochilum mexicanum Burmeister, 1848 Scybalocanthon imitans (Harold, 1868) Deltochilum orbiculare Lansberge, 1874 Scybalocanthon moniliatus (Bates, 1887) Deltochilum peruanum Paulian, 1939 Scybalocanthon pinopterus (Kirsch, 1873) Deltochilum pretiosum Harold, 1875 Scybalocanthon sexspilotus (Gue´rin-Meńeville, Deltochilum pseudoicarus Balthasar, 1939 1855) Deltochilum robustus Molano and Gonzalez, 2009 Scybalocanthon trimaculatus (Schmidt, 1922) Deltochilum tessellatum Bates, 1870 Scybalocanthon zischkai Martıńez, 1949 Deltochilum valgum Burmeister, 1873 Scybalophagus rugosus (Blanchard, 1846) Deltochilum sp. aff. valgum Burmeister, 1873 Streblopus opatroides Lansberge, 1874 Malagoniella astyanax (Olivier, 1789) Streblopus punctatus Balthasar, 1938* Malagoniella cupreicollis Waterhouse, 1890* Sylvicanthon bridarollii (Martıńez, 1949) Pseudocanthon felix (Arrow, 1913) Sylvicanthon furvus (Schmidt, 1921) Pseudocanthon xanthurum (Blanchard, 1843)

Dichotomiini Anomiopus andrei Canhedo, 2006 Canthidium sp. aff. centrale Boucomont, 1928 Anomiopus batesi (Waterhouse, 1891) Canthidium coerulescens Balthasar, 1939 Anomiopus brevipes (Waterhouse, 1891) Canthidium cupreum (Blanchard, 1846) Anomiopus cambeforti Canhedo, 2006* Canthidium sp. aff. deyrollei Harold, 1867 Anomiopus foveicollis Canhedo, 2006 Canthidium discolor Harold, 1867 Anomiopus gilli Canhedo, 2006* Canthidium cf. dohrni Harold, 1867 Anomiopus idei Canhedo, 2006* Canthidium escalerai Balthasar, 1939 Anomiopus intermedius (Waterhouse, 1891) Canthidium sp. aff. funebre Balthasar, 1939 Anomiopus pictus (Harold, 1862) Canthidium gerstaeckeri Harold, 1867 Anomiopus pishtaco Edmonds and Figueroa, 2013* Canthidium sp. aff. gigas Balthasar, 1939 Anomiopus pumilius Canhedo, 2006 Canthidium histrio Balthasar, 1939 Anomiopus smaragdinus (Westwood, 1842) Canthidium kirschi Harold, 1875 Anomiopus validus Canhedo, 2006* Canthidium lentum Erichson, 1847 Ateuchus aeneomicans (Harold, 1868) Canthidium miscellum Harold, 1883 Ateuchus cereus (Harold, 1868) Canthidium cf. onitoides (Perty, 1830) Ateuchus columbianus (Harold, 1868) Canthidium sp. aff. quadridens Harold, 1867 Ateuchus connexus (Harold, 1868) Canthidium cf. ruficolle (Germar, 1824) Ateuchus sp. aff. laevicollis (Harold, 1868) Canthidium thalassinum (Erichson, 1847) Ateuchus peruanus (Balthasar, 1939)* Dichotomius adrastus (Harold, 1875) * Ateuchus pygidialis (Harold, 1868) Dichotomius apicalis (Lu¨derwaldt, 1931) Ateuchus sp. aff. pygidialis Harold, 1868 Dichotomius batesi (Harold, 1869) Ateuchus scatimoides (Balthasar, 1939) Dichotomius belus (Harold, 1869) * Ateuchus sp. aff. setulosus Balthasar, 1939 Dichotomius bicornis (Waterhouse, 1891)* Ateuchus simplex (Le Peletier and Serville, 1828) Dichotomius bicuspis Germar, 1824 Ateuchus striatulus (Preudhomme de Borre, 1886) Dichotomius calcaratus (Arrow, 1913) Ateuchus substriatus (Harold, 1868) Dichotomius camargoi (Martıńez, 1955) Ateuchus viridimicans (Boucomont, 1935) Dichotomius conicollis (Blanchard, 1843) Besourenga horacioi (Martıńez, 1967) Dichotomius cotopaxi (Guerin, 1855) Bdelyrus cochabambae Cook, 2000 Dichotomius cuprinus (Felsche, 1901)* Bdelyrus howdeni Cook, 1998 Dichotomius diabolicus (Harold, 1875) Bdelyrus iquitosensis Cook, 2000* Dichotomius fissus (Harold, 1867) Bdelyrus lobatus Cook, 1998* Dichotomius sp. aff. fonsecae (Lu¨derwaldt, 1924) Bdelyrus parvus Cook, 1998 Dichotomius globules (Felsche, 1901) Bdelyrus pecki Cook, 1998 Dichotomius inachus (Erichson, 1847) Bdelyrus peruviensis Cook, 1998* Dichotomius lucasi (Harold, 1869) Bradypodidium adisi (Ratcliffe, 1980) Dichotomius mamillatus (Felsche, 1901) Canthidium angusticeps (Bates, 1887) Dichotomius melzeri (Lu¨derwaldt, 1922) Canthidium atramentarium Balthasar, 1939 Dichotomius ocellapunctatus (Felsche, 1901) Canthidium basipunctatum Balthasar, 1939 Dichotomius ohausi (Lu¨derwaldt, 1924) Canthidium batesi Harold, 1867 Dichotomius planicollis (Gillet, 1911) Canthidium bicolor Boucomont, 1928 Dichotomius prietoi Martıńez and Martıńez, 1982 VOLUME 88, ISSUE 2 197

Dichotomius problematicus (Lu¨derwaldt, 1924) Ontherus obliquus Geńier, 1996 Dichotomius protectus (Harold, 1867) Ontherus pubens Geńier, 1996 Dichotomius pullus (Felsche, 1910)* Ontherus raptor Geńier, 1996 Dichotomius quinquelobatus (Felsche, 1901) Ontherus rectus Geńier, 1996* Dichotomius robustus (Lu¨derwaldt, 1935) Ontherus sulcator (Fabricius, 1775) Dichotomius satanas (Harold, 1867) Ontherus tenuistriatus Geńier, 1996* Dichotomius semiaeneus (Germar, 1824) Ontherus ulcopygus Geńier, 1996 Dichotomius simplicicornis (Lu¨derwaldt, 1924) Scatimus cucullatus Erichson, 1847* Dichotomius virescens (Lu¨derwaldt, 1924) Scatimus monstrosus Balthasar, 1939 Dichotomius worontzowi (Pereira, 1942) Scatimus sp. aff. onorei Geńier and Kohlmann, Genieridium cryptops (Arrow, 1913 2003 Homocopris torulosus (Eschscholtz, 1822) Scatimus quadricuspis Geńier and Kohlmann, Onoreidium cristatum (Arrow, 1931) 2003* Ontherus alexis (Blanchard, 1846) Scatimus strandi Balthasar, 1939 Ontherus aphodioides Burmeister, 1847 Sinapisoma minuta Boucomont, 1928 Ontherus ashei Geńier, 1996* Trichillum externepunctatum (Preudhomme de Ontherus azteca Harold, 1869 Borre, 1880) Ontherus brevipennis Harold, 1867 Uroxys bahianus Boucomont, 1927 Ontherus bridgesi Waterhouse, 1891 Uroxys elongatus Harold, 1868 Ontherus sp. aff. bridgesi Waterhouse, 1891 Uroxys sp. aff. kratochvili Balthasar, 1947 Ontherus carinifrons Lu¨derwaldt, 1930 Uroxys sp. aff. minutus Harold, 1868 Ontherus edentulus Geńier, 1996 Uroxys peruanus Balthasar, 1940* Ontherus howdeni Geńier, 1996 Uroxys sp. aff. simplex Waterhouse, 1891 Ontherus laminifer Balthasar, 1938 Uroxys variabilis Robinson, 1951

Phanaeini Coprophanaeus callegarii Arnaud, 2002* Megatharsis buckleyi Waterhouse, 1891 Coprophanaeus degallieri Arnaud, 1997 Oruscatus davus (Erichson, 1847) Coprophanaeus ignecinctus (Felsche, 1909) Oxysternon conspicillatum (Weber, 1801) Coprophanaeus lancifer (L., 1767) Oxysternon lautum (MacLeay, 1819) Coprophanaeus larseni Arnaud, 2002 Oxysternon silenus Laporte, 1840 Coprophanaeus ohausi (Felsche, 1911) Oxysternon spiniferum Laporte, 1840 Coprophanaeus parvulus (Olsoufieff, 1924) Phanaeus achilles Boheman, 1858 Coprophanaeus suredai Arnaud, 1996 Phanaeus bispinus Bates, 1868 Coprophanaeus telamon (Erichson, 1847) Phanaeus cambeforti Arnaud, 1982 Dendropaemon angustipennis Harold, 1869 Phanaeus chalcomelas (Perty, 1830) Dendropaemon telephus Waterhouse, 1891 Phanaeus haroldi Kirsch, 1871 Diabroctis mimas (L., 1758) Phanaeus lecourti Arnaud, 2000 Gromphas aeruginosa (Perty, 1830) Phanaeus lunaris Taschenberg, 1870 Gromphas amazonica (Bates, 1870) Phanaeus meleagris Blanchard, 1843 Phanaeus sororibispinus Edmonds and Zidek, 2012 Sulcophanaeus actaeon (Erichson, 1847)* Sulcophanaeus faunus (Fabricius, 1775) Tetramereia convexa (Harold, 1869)

ORPHNINAE Aegidinus petrovi Colby, 2009* Paraegidium costalimai Vulcano, Pereira, and Aegidinus teamscaraborum Colby, 2009 Martı´nez, 1966

ALLIDIOSTOMATINAE Allidiostoma simplicifrons (Fairmaire, 1885) Parallidiostoma tricornum Ocampo and Colby, 2009*

MELOLONTHINAE

Sericini Astaena andicola Frey, 1973* Astaena exquisita Frey, 1973* Astaena biciliata Saylor, 1946* Astaena glabroclypealis Frey, 1974 198 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Astaena moseri Frey, 1973* Astaena pygidia Saylor, 1946* Astaena negligens Frey, 1973* Astaena setosa Frey, 1973* Astaena penai Frey, 1973* Astaena tridentata Erichson, 1847 Astaena peruana Moser, 1918* Raysymmela huanuca Saylor, 1947* Astaena peruensis Frey, 1973* Symmela varians Erichson, 1847* Astaena pottsi Saylor, 1946*

Diplotaxini Liogenys leechi Frey, 1967*

Melolonthini Phyllophaga austera (Erichson, 1847)* Phyllophaga pachypyga (Burmeister, 1855) Phyllophaga jumberea Saylor, 1942* Phyllophaga peruana (Moser, 1918) Phyllophaga marcapatana Moser, 1918* Phyllophaga umbrosa (Erichson, 1847)*

Macrodactylini Ancistrosoma hilare Arrow, 1913 Isonychus cervinalis Frey, 1965* Ancistrosoma intermedium Arrow, 1913* Isonychus cervinoides Frey, 1965* Ancistrosoma klugii Curtis, 1836* Isonychus cervinus Erichson, 1847 Ancistrosoma reductum Frey, 1964* Isonychus egregius Frey, 1965* Ancistrosoma vittigerum Erichson, 1847 Isonychus fraudulentus Frey, 1969* Barybus compacta (Erichson, 1847)* Isonychus nitens Moser, 1921* Barybus peruana Moser, 1918* Isonychus nubeculus Frey, 1969* Barybus squamiger Frey, 1967* Isonychus ovinus Erichson, 1847* Calodactylus abendrothii Kirsch, 1873* Isonychus pavonii Erichson, 1847* Calodactylus heterosquamosus Frey, 1973* Isonychus peruanus Moser, 1921* Ceraspis innotata (Blanchard, 1850)* Isonychus pulchellus Moser, 1918* Ceraspis penai Frey, 1964* Isonychus rosettae Frey, 1969* Ceraspis rubiginosa (Latreille, 1811)* Isonychus saylori Frey, 1969* Ceraspis rufoscutellata Moser, 1919* Isonychus similis Frey, 1973* Ceraspis squamulifera (Moser, 1919) Isonychus simulator Frey, 1969* Chariodactylus sublaevicollis Moser, 1919* Macrodactylus bilineolatus Moser, 1919* Clavipalpus peruanus Moser, 1918* Macrodactylus bistriatus Moser, 1919* Clavipalpus spadiceus (Burmeister, 1855)* Macrodactylus brenskei Moser, 1918* Ctilocephala asperula (Perty, 1830) Macrodactylus cinereus Blanchard, 1850* Dicrania peruana Frey, 1972* Macrodactylus discipennis Moser, 1918* Isonychus arbusticola Erichson, 1847* Macrodactylus marginicollis Moser, 1919* Isonychus cervicapra Frey, 1965* Macrodactylus peruanus Moser, 1919* Macrodactylus sapphirinus Moser, 1919* Plectris molesta (Kirsch, 1873)* Macrodactylus sulcicollis Moser, 1919* Plectris penaella Frey, 1967* Macrodactylus vittipennis Moser, 1919* Plectris penai Frey, 1967* Plectris aberrans Frey, 1964* Plectris sculptipennis Frey, 1974* Plectris candezei Frey, 1967 Plectris tenebrosa Frey, 1967* Plectris kochi Frey, 1967 * Plectris tolimana (Moser, 1921) Plectris lanata Frey, 1964* Plectris tuberculata (Moser, 1919)

Pachydemini Diaphylla hispida Erichson, 1847* Leuretra pectoralis Erichson, 1847

RUTELINAE

Anatistini Spinochlamys macropus (Benderitter, 1921)* Spodochlamys latipes Arrow, 1946 Spodochlamys feyeri Ohaus, 1908 Spodochlamys peruvianus Soula, 2010* Spodochlamys iheringi Ohaus, 1905 VOLUME 88, ISSUE 2 199

Rutelini Acraspedon bernierei Soula, 2002* Lagochile delassisei Soula, 2005* Acraspedon peruvianus Soula, 2002* Lagochile ebrardi Soula, 2010* Aequatoria cludtsi Soula, 2002* Lagochile fuscoviridis Bouchard and Soula, 2005 Aequatoria davidi Soula, 2005* Lagochile peruana peruana Ohaus, 1898 Aequatoria lequericai Soula, 2006* Lagochile peruana huallagensis Soula, 2005* Anticheiroides adamsii nevinsoni (Fowler, 1906)* Lagochile peruana occidentalis Soula, 2005* Anticheiroides davidi Soula, 2006 Lagochile pottgensi Demez and Soula, 2010 Catoclastus chevrolatii Solier, 1851* Lagochile rodriguezi Soula, 2009* Catoclastus jaumesi Soula, 2010* Lagochile santacruzis santacruzis (Machatschke, Catoclastus rabinovichi Martı´nez, 1971* 1972)* Chlorota callegariorum Soula, 2005* Lagochile santacruzis chanchomayoensis Soula, Chlorota chaparroi Curoe and Soula, 2005* 2005* Chlorota chavezlopezi Soula, 2006* Lagochile solimoensis solimoensis Ohaus, 1903 Chlorota columbica columbica (Ohaus, 1912) Lagochile solimoensis oberthuri Soula, 2005 Chlorota columbica peruana Soula, 2002* Lagochile solimoensis wadai Soula, 2005* Chlorota nasuta Ohaus, 1905 Lagochile tibialis (Ohaus, 1935)* Chlorota sergiocastroi Soula, 2008* Lagochile trigona trigona (Herbst, 1790) Chlorota surinama surinama (Ohaus, 1898) Lagochile trigona mancocapaci Soula, 2005* Chlorota surinama iquitosensis Soula, 2005* Lagochile trigona pozuzoensis Soula, 2005* Chrysina argenteola (Bates, 1888) Lagochile vergaracobianae Soula and Curoe, 2005* Chrysophora chrysochlora (Latreille, 1811) Lagochile vasseli Soula, 2010* Cnemida retusa (Fabricius, 1801) Lagochile villatoroae villatoroae Soula, 2005* Dorysthetus andicola Ohaus, 1905 Lagochile villatoroae tingomariaensis Soula, 2005* Dorysthetus fulgida (Waterhouse, 1881) Lasiocala detingomaria Soula, 2006* Dorysthetus mezai Soula, 2005* Lasiocala dioni Soula, 2006* Dorysthetus peruanus (Ohaus, 1905) Lasiocala jenseni Soula, 2006* Epichalcoplethis benjamini Bouchard and Soula, Lasiocala josei Soula, 2006* 2006 Lasiocala lamasi Soula, 2006* Epichalcoplethis gilleti Soula, 2010 Lasiocala schmiti Soula, 2006* Epichalcoplethis santistebani Bouchard and Soula, Lasiocala vasseli Soula, 2010* 2006* Macraspis andicola Burmeister, 1844 Epichalcoplethis schiffleri Bouchard and Soula, 2006* Macraspis assimilis Ohaus, 1908* Eremophygus philippi Ohaus, 1910 Macraspis bicincta bicincta Burmeister, 1844 Exanticheira vidua (Ohaus, 1922) Macraspis chalcea Burmeister, 1844 Exothyridium filippii Soula, 2002* Macraspis chloraspis chloraspis Laporte, 1840 Exothyridium mercieri Soula, 2002* Macraspis chloraspis subandina Soula, 1998 Heterochlorota colini Soula, 2008* Macraspis chrysis (L., 1764) Heterochlorota mathildae mathildae (Ohaus, 1908) Macraspis concoloripes ratcliffi (Soula, 2005) Heterochlorota mathildae peruana Soula, 2002* Macraspis festiva Burmeister, 1844 Homonyx maurettei Soula, 2010 * Macraspis maculata maculata Burmeister, 1844 Homonyx peruanus Ohaus, 1913* Macraspis maculicollis Ohaus, 1905* Homonyx zovii Demez and Soula, 2011 Macraspis martinezi martinezi Soula, 2003 Hypaspidius costatus (Burmeister, 1844)* Macraspis melanaria (Blanchard, 1850) Lagochile aequatorialis aequatorialis Ohaus, 1898 Macraspis peruviana Ohaus, 1898* Lagochile aequatorialis raimondii Soula, 2005* Macraspis pseudochrysis Landin, 1956 Lagochile amazona (Thunberg, 1822) Macraspis stirpita Ohaus, 1914* Lagochile andicola andicola Ohaus, 1903 Macraspis testaceipennis Ohaus, 1898 Lagochile andicola condori Soula, 2005* Macraspis willersi Soula, 2010* Lagochile anophrys (Ohaus, 1914) Macraspis xanthosticta Burmeister, 1844 Lagochile brunnea brunnea (Perty, 1830) Mecopelidnota arrowi Bates, 1904 Lagochile brunnea satipoensis Soula, 2005* Mecopelidnota dewynteri Soula, 2008* Lagochile brunnea tenaensis Soula, 2005* Mecopelidnota marxi Soula, 2008 Lagochile brusteli Soula, 2005* Mecopelidnota mezai Soula, 2008* Lagochile cachetica cachetica Ohaus, 1903 Mesomerodon spinipenne Ohaus, 1905 Lagochile cachetica fusciventris Ohaus, 1912* Microrutela campa (Ohaus, 1922) Lagochile cachetica orientalis Soula, 2005 Microrutela ucalayiensis Jameson, 1997 Lagochile ciliata ciliata Ohaus, 1908 Minidorysthetus hoehnei (Ohaus, 1914)* 200 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Minidorysthetus tingomariaensis Soula, 2006* Pseudohypaspidius silvestrei Soula, 2002* Minidorysthetus ucayaliensis Soula, 2006* Pseudomacraspis affinis affinis (Laporte, 1840) Minidorysthetus vandemergheli Soula, 1998* Pseudomacraspis affinis amazonica Soula, 2002 Oryctomorphus maculicollis Gue´rin-Me´neville, 1838 Pseudomacraspis beryllina beryllina (Erichson, 1847)* Parachlorota estebani Demez and Soula, 2010* Pseudothyridium (Pseudothyridium) bouchardi Parachlorota josei Soula, 2005* Soula, 2002* Parachlorota morettoi Soula, 2002* Pseudothyridium buckwaldi (Ohaus, 1912) Paradorysthetus signatipennis signatipennis (Ohaus, Pseudothyridium ericki Soula, 2006* 1908)* Pseudothyridium hirtum (Kirsch, 1870) Paraptenomela amazona tingomariaensis Soula, 2002* Pseudothyridium juanjosei Soula, 2006* Paraptenomela opalescens (Ohaus, 1935)* Pseudothyridium minettii Soula, 2002* Paratelaugis robusta robusta (Kirsch, 1871) Pseudothyridium oblongum oblongum (Ohaus, 1905)* Pelidnota angiae Demez and Soula, 2009* Pseudothyridium quentini Soula, 2002* Pelidnota bondili Soula, 2006* Ptenomela grangesi Soula, 2006* Pelidnota brusteli Soula, 2010* Ptenomela tavakiliani Soula, 2002* Pelidnota chlorana Erichson, 1847 Ptenomela toulgoeti Soula, 2006* Pelidnota dobleri Frey, 1967 Rutela heraldica Perty, 1832 Pelidnota fusciventris fusciventris Ohaus, 1905* Rutela histrio Sahlberg, 1823 Pelidnota halleri Demez and Soula, 2010* Rutela histrioparilis Jameson, 1997 Pelidnota hernanlequericai Soula, 2006* Rutela laeta (Weber, 1801) Pelidnota hoefigi Ohaus, 1912* Rutela lineola (Linnaeus, 1767) Pelidnota incerta Soula, 2006* Rutela tricolorea (Ohaus, 1905) Pelidnota injantepalominoi Demez and Soula, 2010* Sorocha bousqueti Soula, 2006* Pelidnota lacazei Soula, 2010* Sorocha carloti Demez and Soula, 2010* Pelidnota mezai Soula, 2009* Sorocha castroi Soula, 2008* Pelidnota neitamorenoi neitamorenoi (Soula, 2006) Sorocha champenoisi Soula, 2006* Pelidnota neitamorenoi rodriguezdemendozaensis Sorocha chapellei Demez and Soula, 2010* Soula, 2010* Sorocha damasoi Soula, 2006* Pelidnota ohausi ohausi Frey, 1976 Sorocha jeanmaurettei Demez and Soula, 2010* Pelidnota ohausi piurensis (Soula, 2006)* Sorocha lamasi lamasi Soula, 2006* Pelidnota peslieri Soula, 2009* Sorocha lamasi satipoensis Soula, 2006 * Pelidnota polita (Latreille, 1811) Sorocha maylini Soula, 2006* Pelidnota porioni Soula, 2010* Sorocha similis (Ohaus, 1908)* Pelidnota satipoensis Demez and Soula, 2010* Sorocha yelamosi Soula, 2010* Pelidnota schneideri Soula, 2010* Telaugis aenescens aenescens Burmeister, 1844 Pelidnota subandina subandina Ohaus, 1905 Telaugis aenescens subandina Soula, 1998* Pelidnota testaceovirens felipemezai Soula, 2006* Theuremaripa buchei (Soula, 2002)* Pelidnota toulgoeti Soula, 2006* Theuremaripa imitatrix (Ohaus, 1903) Pelidnota uncinata Ohaus, 1930 Theuremaripa meyeri (Soula, 2005)* Pelidnota unicolor unicolor (Drury, 1778) Thyriochlorota lassalei Soula, 2002* Pelidnota unicolor subandina Soula, 2009* Thyriochlorota villosa (Ohaus, 1908)* Pelidnota werneri Soula, 2006* Tipicha champanheti Soula, 2002* Pelidnota zovii Soula, 2010* Tipicha joliveti Soula, 2002* Pseudochlorota peruana Ohaus, 1905* Vayana melzeri melzeri Ohaus, 1928 Pseudohypaspidius antoinei Soula, 1998* Vayana melzeri subandina Soula, 1998*

Anomalini Callistethus aequatorialis huanapensis (Ohaus, Callistethus suratus (Burmeister, 1844)* 1908)* Callistethus tricostatus (Ohaus, 1897) Callistethus antis (Ohaus, 1903)* Paranomala cincta viridicollis Burmeister, 1844 Callistethus cicatricosa (Perty, 1832) Paranomala hylobia (Ohaus, 1897)* Callistethus eckhardti (Ohaus, 1897)* Paranomala inconstans Burmeister, 1844 Callistethus kulzeri (Frey, 1968)* Paranomala undulata peruviana Gue´rin-Me´neville, Callistethus marginatus (Fabricius, 1792) 1838* Callistethus penai (Frey, 1968)* Strigoderma marginata (Olivier, 1789) Callistethus pyritosus (Erichson, 1847)* Strigoderma peruviensis Blanchard, 1850 Callistethus rufomicans (Ohaus, 1897) Strigoderma sulcipennis sumtuosa Burmeister, 1844 VOLUME 88, ISSUE 2 201

Anoplognathini Platycoelia abdominalis Ohaus, 1904* Platycoelia insolita Smith, 2003* Platycoelia aenigma Smith, 2003* Platycoelia kirschi (Ohaus, 1904)* Platycoelia alternans Erichson, 1847 Platycoelia laelaps (Gutie´rrez, 1951)* Platycoelia baessleri (Ohaus, 1904)* Platycoelia lutescens Blanchard, 1851 Platycoelia burmeisteri Arrow, 1899 Platycoelia marginata Burmeister, 1844 Platycoelia convexa Smith, 2003 Platycoelia peruviana Smith, 2003 Platycoelia flavostriata (Latreille, 1813) Platycoelia pomacea Erichson, 1847 Platycoelia gaujoni Ohaus, 1904 Platycoelia prasina Erichson, 1847 Platycoelia helleri (Ohaus, 1904) Platycoelia rufosignata Ohaus, 1904 Platycoelia inca Smith, 2003* Platycoelia selanderi Martı´nez and Martı´nez, 1994 Platycoelia inflata Ohaus, 1904

Geniatini Bolax albopilosa Ohaus, 1917* Leucothyreus lazarus Ohaus, 1918 Bolax andicola Burmeister, 1844* Leucothyreus saparus Ohaus, 193* Bolax boliviensis Ohaus, 1898 Lobogeniates bicolor Ohaus, 1917 Bolax cupreoviridis Ohaus, 1931* Trizogeniates aphilus Villatoro, 2002* Bolax disgamia Ohaus, 1917* Trizogeniates apicalis Ohaus, 1917* Bolax glabripennis Ohaus, 1917* Trizogeniates barrerai Martı´nez, 1965* Bolax gonzalofideli Soula, 2010* Trizogeniates caiporae Villatoro, 2002 Bolax incogitata Dohrn, 1883 Trizogeniates catsus Villotoro, 2002 Bolax malkini Soula, 2010* Trizogeniates crispospinatus Villatoro, 2002* Bolax nigriceps Ohaus, 1917* Trizogeniates laticollis Ohaus, 1931 Bolax robackeri Soula, 2010* Trizogeniates ohausi Villatoro, 2002 Bolax rutila Erichson, 1847* Trizogeniates planipennis Ohaus, 1917 Trizogeniates temporalis Ohaus, 1917 Geniates balzapamae Ohaus, 1917 Trizogeniates tibialis Ohaus, 1917 Leucothyreus baeri Ohaus, 1917* Trizogeniates trivittatus Ohaus, 1917 Leucothyreus demetrius Ohaus, 1918*

DYNASTINAE

Cyclocephalini Acrobolbia macrophylla Ohaus, 2012 Cyclocephala brevis Ho¨hne, 1923 Ancognatha castanea Erichson, 1847 Cyclocephala colasi Endro¨di, 1964 Ancognatha erythrodera (Blanchard, 1841) Cyclocephala confusa Endro¨di, 1966 Ancognatha humeralis Burmeister, 1847 Cyclocephala contracta Kirsch, 1873 Ancognatha lutea Erichson, 1847 Cyclocephala couturieri Dechambre, 1999 Ancognatha scarabaeoides Erichson, 1847 Cyclocephala dilatata (Prell, 1934) Ancognatha vulgaris Arrow, 1911 Cyclocephala diluta Erichson, 1847 Aspidolea brunnea Ho¨hne, 1922 Cyclocephala discolor (Herbst, 1792) Aspidolea collaris Endro¨di, 1965* Cyclocephala dispar (Herbst, 1792) Aspidolea fuliginea (Burmeister, 1847) Cyclocephala flavoscutellaris Ho¨hne, 1923 Cyclocephala flora Arrow, 1911 Aspidolea laticeps Harold, 1869 Cyclocephala freyi Endro¨di, 1964 Aspidolea lindae Ratcliffe, 1978 Cyclocephala fulgurata Burmeister, 1847 Aspidolea mimethes (Ho¨hne, 1922)* Cyclocephala fulvipennis Burmeister, 1847 Aspidolea notaticollis Ho¨hne, 1922 Cyclocephala genieri Joly, 2010* Aspidolea singularis Bates, 1888 Cyclocephala goetzi Endro¨di, 1966* Aspidolea suturalis Ho¨hne, 1922 Cyclocephala guycolasi Dechambre, 1992 Augoderia freyi Endro¨di, 1976* Cyclocephala hirsuta Ho¨hne, 1923 Chalepides paradytis Ponchel and Dechambre, 2003 Cyclocephala inca Endro¨di, 1966 Cyclocephala affinis Endro¨di, 1966 Cyclocephala isabellina Ho¨hne, 1923 Cyclocephala almitana Dechambre, 1992 Cyclocephala kaszabi Endro¨di, 1964 Cyclocephala altamontana Dechambre, 1999 Cyclocephala ligyrina Bates, 1888 Cyclocephala amazona (L., 1767) Cyclocephala lineigera Ho¨hne, 1923 Cyclocephala bicolor Laporte,1840 Cyclocephala liomorpha Arrow, 1911 202 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Cyclocephala lunulata Burmeister, 1847 Cyclocephala spilopyga Erichson, 1847 Cyclocephala macrophylla Erichson, 1847 Cyclocephala stictica Burmeister, 1847 Cyclocephala mannheimsi Endro¨di, 1964 Cyclocephala testacea Burmeister, 1847 Cyclocephala marginalis Kirsch,1870 Cyclocephala tronchonii Martıńez, 1975* Cyclocephala mecynotarsis Ho¨hne, 1923 Cyclocephala tylifera Ho¨hne, 1923 Cyclocephala melanocephala (Fabricius, 1775) Cyclocephala verticalis Burmeister, 1847 Cyclocephala molesta Endro¨di, 1969 Cyclocephala viridis Dechambre, 1982 Cyclocephala moreti Dechambre, 1992 Cyclocephala zurstrasseni Endro¨di, 1964* Cyclocephala morphoidina Prell, 1937 Dyscinetus dubius (Olivier, 1798) Cyclocephala munda Kirsch, 1870 Dyscinetus dytiscoides (Arrow, 1911) Cyclocephala obscura Endro¨di, 1966* Dyscinetus olivaceus Ho¨hne, 1923 Cyclocephala ocellata Bolı´var y Pieltain, Jimeńez- Dyscinetus paradytis Ponchel and Dechambre, 2003 Asuá, and Martıńez, 1963 Erioscelis peruana Saylor, 1946* Cyclocephala octopunctata Burmeister, 1847 Erioscelis proba (Sharp, 1877) Cyclocephala panthera Dechambre, 1979 Harposcelis paradoxus Burmeister, 1847 Cyclocephala paraflora Martıńez, 1978 Mimeoma signatoides Ho¨hne, 1923 Cyclocephala paraguayensis Arrow, 1903 Stenocrates bicarinatus Robinson, 1948 Cyclocephala peruana Endro¨di, 1966* Stenocrates carbo Prell, 1937 Cyclocephala pilosa Dupuis, 2006* Stenocrates celatus Prell, 1937 Cyclocephala prolongata Arrow, 1902 Stenocrates clipeatus Endro¨di, 1966 Cyclocephala pugnax Arrow, 1914 Stenocrates cognatus Endro¨di, 1966 Cyclocephala quadripunctata Ho¨hne, 1923 Stenocrates cultor Burmeister, 1847 Cyclocephala rufovaria Arrow, 1911 Stenocrates haackae Ratcliffe, 1979 Cyclocephala rustica municipalis Ho¨hne, 1923 Stenocrates holomelanus (Germar, 1824) Cyclocephala saltini Ratcliffe, 2008 Stenocrates minutus Endro¨di, 1966 Cyclocephala scarabaeina (Gyllenhal, 1817) Stenocrates nasutus Dechambre, 1979* Cyclocephala sexpunctata Laporte, 1840 Stenocrates popei Endro¨di, 1971 Cyclocephala simulatrix Ho¨hne, 1923

Pentodontini Bothynus entellus (LePeletier and Serville, 1828) Oxyligyrus peruanus Endro¨di, 1966 Bothynus lancifer Dechambre, 1981* Parapucaya amazonica Prell, 1934 Diloboderus abderus (Sturm, 1826) Parapucaya nodicollis (Kirsch, 1873) Heteroglobus obesus Dupuis and Dechambre, 2008* Pentodina peruviana Endro¨di, 1968 Hylobothynus obesus Ohaus, 1910 Piscoperus paracanicola Ratcliffe and Giraldo, 2014* Oxyligyrus contractus Dupuis, 2010* Tomarus bituberculatus (Palisot de Beauvois, 1805) Tomarus burmeisteri (Steinheil, 1872) Tomarus maternus (Prell, 1937) Tomarus ebenus (DeGeer, 1774) Tomarus peruvianus (Endro¨di, 1970) Tomarus gyas Erichson, 1847 Tomarus similis (Endro¨di, 1968) Tomarus maimon Erichson, 1847 Tomarus villosus (Burmeister, 1847)

Oryctini Coelosis biloba (L., 1767) Heterogomphus ulysses Burmeister, 1847 Enema pan (Fabricius, 1775) Megaceras brevis Dechambre, 1999 Heteroglobus obesus Dupuis and Dechambre, 2008 Megaceras briansaltini Ratcliffe, 2007* Heterogomphus arrowi Prell, 1912* Megaceras endroedii Dechambre, 1998 Heterogomphus dilaticollis Burmeister, 1847 Megaceras inflatum Prell, 1934 Heterogomphus hirticollis Prell, 1912* Megaceras laevipenne Prell, 1914 Heterogomphus hirtus Prell, 1912 Megaceras morpheus Burmeister, 1847 Heterogomphus incornutus Prell, 1912* Megaceras philoctetes (Olivier, 1789) Heterogomphus mirabilis Prell, 1912* Megaceras porioni Dechambre, 1981 Heterogomphus ochoai Martı´nez, 1966 Megaceras quadraticollis Dechambre, 1975 Heterogomphus orsilochus Erichson, 1847 Podischnus oberthueri Sternberg, 1907 Heterogomphus peruanus Endro¨di, 1976* Podischnus sexdentatus (Taschenberg, 1870) Heterogomphus pilosus Dechambre, 1998 Strategus aloeus (L., 1758) Heterogomphus porioni Dechambre, 1998 Strategus jugurtha Burmeister, 1847 Heterogomphus rugicollis Prell, 1912 Strategus surinamensis hirtus Sternberg, 1910 Heterogomphus rubripennis Prell, 1912 VOLUME 88, ISSUE 2 203

Phileurini Amblyodus castroi Grossi and Grossi, 2011 Homophileurus waldenfelsi Endro¨di, 1978 Amblyoproctus chalumeaui Endro¨di, 1977 Microphileurus caviceps Kolbe, 1910 Amblyoproctus piliger (Perty, 1830) Microphileurus subulo Prell, 1912* Amblyoproctus rugosus (Erichson, 1847) Oryctophileurus armicollis Prell, 1911 Archophileurus aper Endro¨di, 1977 Palaeophileurus carbo Ratcliffe, 2002 Archophileurus burmeisteri (Arrow, 1908) Palaeophileurus erebus Ratcliffe, 2002 Archophileurus oedipus (Prell 1912) Palaeophileurus marcusoni Ratcliffe, 1998 Archophileurus peruanus Endro¨di, 1977 Palaeophileurus ocampoi Neita and Ratcliffe, Archophileurus sus Dechambre, 2006* 2012 Hemiphileurus brasiliensis Endro¨di, 1978 Palaeophileurus proximus Dechambre, 1997 Hemiphileurus depressus (Fabricius, 1801) Palaeophileurus sclateri (Bates, 1887) Hemiphileurus elongatus Dupuis and Dechambre, Phileucourtus bicornutus Dechambre, 2008* 2000* Phileurus angustatus Kolbe, 1910 Hemiphileurus howdeni Endro¨di, 1978* Phileurus didymus (L., 1758) Hemiphileurus isabellae Dupuis, 2004* Phileurus excavatus Prell, 1911 Hemiphileurus kahni Dupuis and Dechambre, Phileurus kaszabi Endro¨di, 1978 2000* Phileurus valgus (Olivier, 1789) Homophileurus quadrituberculatus (Palisot de Beauvois, 1805)

Agaocephalini Aegopsis chaminadei Dechambre, 2000 Brachysiderus quadrimaculatus Waterhouse, 1881 Aegopsis peruvianus Arrow, 1941* Mitracephala humboldti Thomson, 1859

Dynastini Dynastes hercules (L., 1758) Golofa eacus Burmeister, 1847 Dynastes neptunus (Quensel, 1806) Golofa spatha Dechambre, 1989 Golofa aegeon (Drury, 1773) Golofa testudinarius (Prell, 1934)* Golofa clavigera (L., 1771) Golofa unicolor (Bates, 1891)

CETONIINAE

Cetoniini Euphoria steinheili Janson, 1878

Gymnetini Desicasta lobata (Olivier, 1789) Gymnetis pantherina (Blanchard, 1843) Desicasta purpurascens (Schoch, 1898) Gymnetis rufilateris (Illiger, 1800) Gymnetis balzarica Janson, 1880 Gymnetis subpunctata Westwood, 1874 (5? G. varia- Gymnetis coturnix Burmeister, 1842 (5? G. phasia- bilis Moser, 1921) nus Burmeister, 1842) Hoplopyga liturata (Olivier, 1789) Gymnetis mathani Pouillaude, 1913 Hoplopyga peruana Moser, 1912 Gymnetis holoserica Voet, 1778 (5? G. chancha- Marmarina maculosa (Olivier, 1789) mayensis Pouillaude, 1913) Gymnetis pardalis (Gory and Percheron, 1833) (5? G. cupriventris Janson, 1880)

Cremastocheilini Cyclidius lacordairei Thomson, 1860* Genuchinus sp.

Trichiini Golinca davisii Waterhouse, 1877* Golinca ishiharai Nagai, 1994* 204 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Acknowledgments We are grateful to Caroline Chaboo (University of Kansas) for inviting us to participate in the “Beetles of Peru” project and affording us the opportunity for advancing scarab research in Peru. We thank Trond Larsen (Conservation International, Arlington, VA) for sharing his records of Scarabaeinae from Peru. This work was supported, in part, by a Biotic Surveys and Inventory Grant from the National Science Foundation (DEB-0716899) to B. C. Ratcliffe and R. D. Cave. We acknowledge NSF-EPSCoR #66928 (PI: CS Chaboo) and the University of Kansas’ Department of Ecology and Evolutionary Biology-General Research Fund (PI: CS Chaboo) for supporting the “Beetles of Peru” project.

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